EPA-600/3-77-108
September 1977
Ecological Research Series
       A FLOW-THROUGH TESTING PROCEDURE  WITH
                          DUCKWEED, (lemna minor  L)
                                            imental Research Laboratory
                                        Office of Research and Development
                                       U.S. Environmental Protection Agency
                                              Duluth, Minnesota 55804

-------
                 RESEARCH REPORTING SERIES

Research reports of the Office of Research and Development, U.S. Environmental
Protection Agency, have been grouped into nine series. These nine broad cate-
gories were established to facilitate further development and application of en-
vironmental technology. Elimination  of  traditional grouping was consciously
planned to foster technology transfer and a maximum interface in related fields.
The nine series are:

      1.   Environmental Health Effects Research
      2.   Environmental Protection Technology
      3.   Ecological Research
      4.   Environmental Monitoring
      5.   Socioeconomic Environmental Studies
      6.   Scientific and Technical Assessment Reports (STAR)
      7.   Interagency Energy-Environment Research and Development
      8.   "Special"  Reports
      9.   Miscellaneous Reports

This report has been assigned to the ECOLOGICAL RESEARCH series. This series
describes research on the effects of pollution on  humans, plant and animal spe-
cies, and materials.  Problems are assessed for their long- and short-term influ-
ences. Investigations include formation, transport, and pathway studies to deter-
mine the fate of pollutants and their effects. This work provides the technical basis
for setting standards to minimize undesirable changes in living organisms in the
aquatic, terrestrial, and atmospheric environments.
This document is available to the public through the National Technical Informa-
tion Service. Springfield, Virginia 22161.

-------
                                                       EPA-600/3-77-108
                                                       September 1977
A FLOW-THROUGH TESTING PROCEDURE WITH DUCKWEED  (LEMNA MINOR L.)
                               by

                     Charles T. Walbridge
           Environmental Research Laboratory-Duluth
                   Duluth, Minnesota  55804
           ENVIRONMENTAL RESEARCH LABORATORY-DULUTH
             OFFICE OF RESEARCH AND DEVELOPMENT
             U.S. ENVIRONMENTAL PROTECTION AGENCY
                   DULUTH, MINNESOTA  55804

-------
                                  DISCLAIMER
     This report has been  reviewed  by  the Environmental  Research Laboratory-
Duluth, U.S. Environmental Protection  Agency,  and approved for  publication.
Mention of trade names  or  commercial products  does not constitute endorsement
or recommendation  for use.
                                      ii

-------
                                  FOREWORD
     Our nation's fresh waters are vital  for  all  animals  and  plants,  yet our
diverse uses of water —  for recreation,  food,  energy,  transportation,  and
industry — physically and chemically  alter lakes,  rivers,  and  streams.   Such
alterations threaten terrestrial organisms, as  well  as  those  living  in  water.
The Environmental Research Laboratory  in  Duluth,  Minnesota, develops  methods,
conducts laboratory and field studies, and extrapolates research findings

     —to determine how physical and chemical pollution affects aquatic
       life,

     —to assess the effects of ecosystems on pollutants,

     —to predict effects of pollutants on large  lakes  through  use of
       models, and

     —to measure bioaccumulation of pollutants in  aquatic  organisms  that
       are consumed by other animals,  including man.

     This report describes a flow-through testing procedure for duckweed.
It identifies the variables that have  to  be controlled  and  the  ranges to
which they should be confined.  Copper sulfate  is used  as an  example  toxicant.
                                      Donald  I. Mount,  Ph.D.
                                      Director
                                      Environmental  Research  Laboratory
                                      Duluth, Minnesota
                                      iii

-------
                                 ABSTRACT
     Lemna minor is one of the smallest flowering plants.  Because of its
floating habit, ease of culture, and small size it is well adapted for
laboratory investigations.  Procedures for flow-through tests were developed.
Testing procedures were developed with this apparatus.  By using the techniques
described here, the effects of nutrients or toxicants, singly or in combination,
can be determined in several concentrations with several replicates of each.
Responses which can be measured include changes in growth rate, changes in
death rate, changes in timing of division of colonies, color changes, changes
in variability, and modification of the flowering response.  Emphasis here is
on changes in growth rate determined either by daily frond counts or by final
frond numbers.
                                      IV

-------
                                  CONTENTS
Foreword	iii
Abstract	iv
Acknowledgments	vi

     1.  Introduction  	   1
     2.  Conclusions and Recommendations 	   3
     3.  Methods	   4
              Physical methods 	   4
              Biological methods 	   /
     4.  Results	   9
     5.  Discussion	13

Recommended Methods  	  15
References	17
Bibliography	19

-------
                                ACKNOWLEDGMENTS
     The author  gratefully  acknowledges  the technical guidance and editorial
assistance of Dr. Richard L.  Anderson of the Environmental Research Laboratory-
Duluth.
                                      vi

-------
                                   SECTION  1

                                 INTRODUCTION
     Lemna minor L., lesser duckweed,  is  common  throughout the world.   In
fact, it is absent only  from  the polar regions and  a few parts of the  tropics.
Lemna minor is a small floating plant  consisting of a single leaf-like frond
that is flat, more or less oval in  outline,  and  1.5 - 4 mm long.   A single
root is attached about a third of the  way from the  narrow end of  the frond.
The root, which is terminated by a  prominent calyptra,  or root cap,  may be as
long as 10 cm.  Vegetative reproduction is accomplished by the growth  of
daughter fronds alternately from a  pair of reproductive pouches at the sides
of the root-bearing end  of the frond.   Flowering is rare, but the flowers also
arise from the reproductive pockets.   A Lemna colony is made up of several
generations of daughter  fronds still attached to the parent.  In  its natural
environment ]j. minor is  eaten heavily  by  some wildfowl, beaver, and  muskrat.
It serves as physical support for epiphytic  organisms,  which are  in  turn
eaten by aquatic crustaceans, molluscs, annelids, and insect larvae.  For fish
L_. minor provides food,  shelter, and shade,  though  it tends to shade out
other aquatic plants that might produce food more efficiently (Muenscher 1944,
Sculthorpe 1967).

     The Lemnaceae in general have  a number  of characteristics that  make them
useful laboratory organisms,  such as ease in obtaining  clonal material,
simple methods of axenic culture, and  small  size.   A consequence  of  this
combination of properties is  that a large background of research  has been
developed for this species.

     Static tests are conducted in  a non-renewed solution of the  material under
investigation.  Also, the plants may be transferred into fresh solutions during
the test.  Several authors describe applicable techniques.  Fromm (1946, 1951,
1960) used greenhouse lighting and  20  plants per flask with four  flasks per
concentration.  Mitchell et al. (1958) recommended  sterilization  of  the
plants and culture medium for some  uses.   Further,  they control light  (700 -
1000 ft-C for 12 hrs per day) and temperature (24°- 27°C).  Blackman (1952)
controlled light and temperature.   He  did not use sterile solutions  or
sterilize the plants, but instead changed solutions every 48 hrs  to  keep
contamination down as well as to keep  the pH and the chemical concentrations
within reasonable ranges.  Fekete ££ aJ^-  (1976)  used a bioassay procedure to
evaluate the release of  available phosphorus from pond  sediments.  Here again,
greenhouse lighting was  used.

     Advantages of the static and renewal procedures lie in the small  quantities
of solution required and in the simplicity of the experimental design; no
delivery system is required.  The major limitation  of a static experiment is

-------
that the concentrations  of  active materials in the experimental container may
change over  the  test  interval  through degradation, precipitation, etc.

     Flow-through  and recirculating systems of varying degrees of sophistication
have been designed.   These  systems range from single open chambers, through
which the solution moves by a  siphon arrangement (Mendiola 1919), to complex
aseptic systems  in which the area of a raft of plants can be kept constant so
that the effects of  crowding can be eliminated (Erismann and Finger 1968) or
systems in which samples may be taken automatically without violating aspesis
(Strasser 1971).

     Ashby ^t  al.  (1928).designed a recirculating system that was not aseptic.
Eichorn and  Augsten  (1969)  cultured Wolffia arrhiza in a chemostat under
continuous illumination.

     The Lemnaceae have  also been used in.systems for testing the effects of
various air  pollutants on plants.  Todd et_ a^. (1956) tested the effects of
air containing ozone  and ozonated hexane.   Feder and Sullivan (1969) studied
the reduction  of frond multiplication and  floral production by ozone.  Scharer
et al. (1975)  showed  that Lemna minor could metabolize sulfur dioxide from
the air if it  was  made available in sublethal concentrations.

     The cultivation  and testing of Lemna  minor have produced a number of
ways to quantify responses. These depend  on such things as frond death,
premature division of colonies, alteration of flowering response, and dry
weight changes.  At  least two  methods of visually evaluating frond condition,
are available.   One  is to assign degree-of-damage numbers to the fronds, e.g.,
from zero (no.damage) to five  (complete loss of green color) (Parker 1965).
The second visual  method involves frond counts, which have been shown to be a
reliable measure of  growth  response in the Lemnaceae.  These counts compare
well with dry  weight  changes and area changes.  Ashby and Oxley (1935) and
Fromm (1960) have  used this technique.

     The genus Lemna  is  unusual in that plants are not damaged by what might
be considered  excessive  day lenghts, including continuous illumination with
no dark period.  The  continuous growth of  Lemna in a 24-h photoperiod is
useful in reducing the duration of the tests being performed (Clark 1925,
Landolt 1957).

     The light intensities  that can be used by Lemna minor range from 1,600 to
17,000 lux,  but  there is no appreciable increase in growth above about 8,000
lux. (Ashby and Oxley  1935). The lower end of this range is easily attainable
in laboratory  situations.

     The procedures  described  above were not designed with simplicity in mind,
so this work was undertaken to provide a fast, simple, and•inexpensive technique
for evaluating stimulatory  or  inhibitory effects of pollutants on a typical
aquatic plant.

-------
                                   SECTION  2

                       CONCLUSIONS AND  RECOMMENDATIONS
     The lesser duckweed  (Lemna minor  L.)  is  a  suitable  higher  plant for
testing in a flow-through system.   The testing  method  developed in this
study should be useful for evaluating  effects of  toxicants,  enrichments, and
otherwise altered conditions on higher aquatic  plants.   A procedure for
using Lemna minor as a test organism is recommended.

-------
                                 SECTION 3

                                  METHODS


Physical Methods

     Lake Superior water, temperature-controlled but otherwise unmodified,
was delivered at regular intervals by use of a delivery chamber that
periodically emptied itself by a self-starting siphon (Hero of Alexandria,
ca. 62 AD).  The periodic flow of water operated a chemical-metering apparatus
designed for highly water-soluble materials (Mount and Brungs 1967) (Figure 1).
This apparatus added a solution of plant nutrients to the feedwater of a
proportional diluter (Mount and Brungs 1967).  This diluter was provided with
a mechanical syringe injector which added copper sulfate when this chemical was
used as a toxicant (DeFoe 1975) .  The diluter delivered the resulting series of
concentrations through glass delivery tubes into battery jars.  The jars had a
capacity of 1.4 1 each.  The delivery tubes were wrapped in black polyethylene
film to reduce growth of attached algae on the inner surfaces.  Toxicant was
added by an injector system designed to add microliter quantities to the water
to operate as the diluter cycled.

    The battery jars drained through a notch cut 4 cm down from the top edge.
A silicone-based glass and ceramic adhesive was used to fasten stainless steel
bolting cloth (screen) across this opening.  This allowed the water to drain
out, but prevented any loss of experimental plants.  The screen used here was
105 mesh with 0.165-am openings.

     Four separate battery jars were used for each concentration of the material
being tested.  Individual colonies of duckweed were kept separated and centered
in the jars by seven-compartment floats (Figure 2).  The separator consisted of
seven rings, 1-cm sections of 2—cm glass tubing; six of these were glued in a
hexagon around the seventh.  This separator was glued between and supported by
two air-filled 4-, dram glass vials with plastic caps.  The float was kept
centered with a hook made of bent stainless steel wire which was hung over the
edge of the jar.  The separation of the colonies made it possible to monitor
each one individually.  An index mark on one float showed the starting point
so that data for each colony could be recorded separately and consistently
(Figure 2).

    Continuous illumination was provided by fluorescent tubes, half of which
were Durotest Optima and the remainder Gro Lux, Wide Spectrum.  The distance
from the fluorescent tubes to the frond level was 20-25 cm.  This provided a
light intensity of 2,700 lux; the standard deviation (s) was 2.50 lux.

     Temperatures in the test chambers were maintained at 22.8 C (s = 0.6 ).
Alkalinity, pH, hardness, and acidity were measured at the beginning and the
end of each experiment (American Public Health Association 1971).

-------
                        LAKE
                        WATER I
     NUTRIENT
     SUPPLY
                                        TO TEST CHAMBERS   ,
                                                                  TOXICANT
                                                                  SYRINGE
Figure 1.  Nutrient- and toxicant-adding diluter.

-------
           Index mark
                            Float
                        Float  in place
Figure 2., Duckweed testing chamber and float.

-------
     The flow rate  into  the  system was  1.2  1 of water (s = 0.13 1) delivered
every 15.6 min  (s = 2.0);  this was mixed  with 2.5 ml  (s = 0.15 ml) of nutrient
solution.

     Aeration of the  test  chambers was  not  necessary.  Lemna minor fronds have
adequate gas exchange  through their dorsal  surfaces.

Biological Methods

     The clone  that was  used (Lemna minor L.  #6)  was  originally collected in
Florida in the  vicinity  of Boca  Raton some  time before  1964.   This clone has
been in culture at  the Environmental Research Laboratory-Duluth since 1971.
Except where otherwise noted, colonies  of two fronds  each were used to
start a test.   One  colony was placed in each retaining  ring of the float.
This resulted in seven colonies  per jar and,  since each concentration was
replicated four times, 28  colonies per  test concentration.

     To reduce  possible  effects  of attached algal growth on surfaces in contact
with the test solution,  on the 4th day  of a 7-day test  the colonies were
transferred into clean floats in clean  battery jars.  The water in the
replacement battery jars was lake water that  had  been allowed to equilibrate
to room temperature for  16-24 hrs.   When  copper toxicant was  used, the water
in the jars was dosed with the appropriate  concentrations of  copper before
the transfer.   Thus the  duckweed was continuously exposed to  the copper
concentrations.  In the  copper experiment the glassware cleaning procedure
included acid washing.

     In- the course  of experimentation two kinds of nutrients  were used.   The
first was a commercial soluble fertilizer (Garden Life,  Science Products
Company, Chicago, Illinois   60614)  with an  analysis 10% nitrogen,  15% phosphorus,
and 14% potassium.  The  stock concentration was 1.24  mg/1.   The second
nutrient was Hutner's Medium (Hutner 1953).

     Stock cultures were maintained continuously  under  the same conditions as
the test organisms, lacking  only the toxicant additions.

     Growth rate (K) was calculated by  the  formula

                          Log1Q    (Fd)  -  Log 1Q  (Fo)
                    K =      ~ ~
where Fo is the original frond number, Fd  is  the  frond number  on  day  d^,  and
d is the number of days involved  (Hillman  1961).  When analysis of  variance
or regression were used, individual frond  counts  were used, not chamber
averages.

     Plant response for the first 2 or 3 days  reflects a  combination  of
experimental and pre-experimental conditions;  therefore,  growth rates from
this period were not used  (Fromm 1960).  Because  it  is difficult  to maintain
identical light intensities at the surface of  the experimental containers

-------
and because the light output of  fluorescent  tubes  changes with  time,  two
experiments were undertaken to determine whether  the  light variations over
the test bench correlated  significantly with frond numbers at termination.

     An experiment was undertaken  to  correlate  the effects of the  differing
flow rates into the  separate containers with the  frond  numbers  at  termination.
A comparison was made between a  series of  static  concentrations of a  commercial
soluble fertilizer  (Garden Life) and  a continuously replaced  solution of  the
same material.

     The purpose of  another experiment was to determine when  reduction of
growth due to crowding within the  individual float chambers became significant.
Chambers were given  six  two-frond  colonies each instead of one.

     Because unequal amounts of  damage may be caused  in handling or in the
reduction of the colonies  to two-frond units, minimally handled colonies
were compared with  colonies that might be  expected to show a  maximum  response
to handling.  The unmodified colonies were simply transferred  intact  from
stock cultures to test chambers.   The modified colonies were  carefully broken
into two-frond units and the roots were  excised.

     The method developed  here was tested  with copper sulfate as the  toxicant.
ED50 used here indicates a reduction  of  the production of new fronds  by half;
this is derived from the difference between the final frond number and the
original frond number.   Because  copper at  31 ppb  produced more  growth than
the control Lake Superior  water  at 1  ppb,  growth reduction was  calculated
from this level as  a starting point.

-------
                                   SECTION  4

                                   RESULTS
Effect of Light Variation

     In two experiments no correlation was  found  between variations in light
intensity at frond level and  frond numbers  at  termination.   The mean of the
first experiment was 2,317 lux  (s =  219)  and of  the  second  experiment, 2,749
lux (s = 221).

Effect of Flow-Rate Variation

     A significant correlation  (0.57) was found  between flow rate and final
frond number in each test chamber (Figure 3).

Effect of Static vs. Flowing  Conditions

     The flowing concentrations of the soluble fertilizer at 1.24 mg/1
produced growth that was not  significantly  different from the static
concentration of the same fertilizer at 100 mg/1  (Figure 4).

Effect of Handling

     Reduction of the colonies  to two-frond, root-excised colonies did not
produce a significant reduction in growth rate.

Effect of Copper Sulfate

     Linear regression was used for  the data from 31 ppb to 210 ppb.  The
7-day ED50 was 119 ppb copper,  the upper  confidence  limit was 151, and the
lower confidence limit was 105  (Figure 5).

Effect of Crowding

     A statistically significant reduction  in  growth was noted when the area
within the individual float chamber  was reduced to 8.0 mm2  per frond.  Three
days before this area was reached the available area was 12.5 mm2 per frond.

-------
       50
     0
     &

     E
     rs
     c
     Q
                    30
                                                       j	1
                          Flow rate (ml/cycle)
75
.Figure 3.  Correlation between  flow rate in chambers and frond numbers at

           end of 7 day test.   Cycle time is 3.84 cycles per hour.
                                   10

-------
        Q>
        E
           12
           10
        o
        Q.

        I  8

        S.

        CD  6
        73

        O
        O  2

        il
N=77
             N=84
N=84
                                     * significantly different
                                       from control
                                      N=77
                                                  N=84
                       control
                     flow-through
                      0.001 g/l
             static
             1.0 g/l
static
O.I g/l
 static
0.01 g/l
  static
0.001 g/l
Figure 4.  Growth differences between flowing and static conditions at various nutrient concentrations.

-------
 o
 e

  "4
 c
 o
 o
 0)
    0
ED50
                  10      20     40        100    200 300  500

                            Copper (ppb)
Figure 5.  Growth responses to copper, as copper sulfate.
                                12

-------
                                  SECTION 5

                                  DISCUSSION

     These experiments were undertaken in an attempt  to  develop  a  flow-
through testing procedure with duckweed as the  test organism.  The method
was produced by combining earlier work in this  area with simplifications
derived from experiments.  The goal was to create  fast,  simple,  and inexpensive
techniques for evaluating stimulatory or inhibitory effects  of various
pollutants.  Where comparison is possible, the  results produced  here are
consistent with those reported earlier.  However,  in  much of the research  with
duckweed one or more of the important variables  are poorly controlled.

     Light output of the fluorescent tubes can  vary over a fairly  wide range,
both in space (across the table) and time  (between experiments), so it
becomes necessary to monitor these changes and  periodically  determine whether
there are significant correlations between light intensities and growth rates
over the intensity range utilized.  The combination of fluorescent tubes used
here (50:50 Gro Lux Wide Spectrum and Duro Test  Vita  Light)  is standard
practice at the Environmental Research Laboratory-Duluth.  The industry-derived
Color Rendering Index (CRI) for each of these is 55 and  92,  respectively.
Daylight is assigned a CRI of 100.  If a better  approximation of daylight  is
needed, only tubes having a CRI of greater than  90 should be used.

     Flow rates vary among chambers, and growth  rates can vary significantly
with them.  This flow variation should be kept  to  a minimum, and the
correlation between flow rate and growth rate should  be  determined periodically.

     Flowing conditions require far less in the  way of nutrient  concentration.
This can have the effect of reducing the amount  of interaction between  the
necessary nutrients and the material being tested.

     The reduction of the colonies to a standardized  colony  (two fronds, roots
exicised) causes no reduction in growth rate.   It  then is reasonable to start
with such a standard colony for reasons of uniformity and because  the
statistical treatments are somewhat simplified.

     In the test of copper sulfate toxicity, which served here only as  a
test of the procedure, it should be noted  that  the 7-day ED50 of 119 ppb
copper indicates only part of the copper toxicity  problem with duckweed.
Copper is one of several metals that bioaccumulate in plants so  that subsequent
complications are to be expected.
                                      13

-------
     The effects of crowding were apparent at 8.0 mm2 per frond,  but because
the response of duckweed takes an area of 2-3 days to appear,  12.5 mm2,  the
area per frond 3 days earlier, is taken as the upper limit of  allowable
crowding.

     This system or a similar one may be adapted to study the  effects of air
pollution on plants.
                                      14

-------
                              RECOMMENDED  METHODS


     The following recommended  experimental  conditions and procedures are
based on the results of  the  experiments described.

     1)  Initial conditions  and measurements:

          a)  Determine  whether the variations  in growth  are  correlated  with
variations in light intensity or  flow rate.   In any  case,  light  intensity
and flow rate should be  as constant as possible.  Because  changes  in light
intensity and flow rate  are  relatively slow, measurements  may be made between.
not during, experiments.

          b)  For starting the  test and replacing the  glassware, use water
of the same temperature  and  the same concentration of  toxicants  as that  from
which the plants are being transferred.

          c)  The stock  cultures  of experimental plants should be  maintained
in a situation as close  to the  experimental  conditions as  possible.   A two-frond,
root-excised standard colony  reduces transfer of epiphytic algae and makes
growth rates of colonies directly comparable.

          d)  A proportional  diluter is not  necessarily the only way to  provide
the toxicant solutions,  but  any system used  should have fail-safe  operation,
so that if water flow stops,  toxicant flow stops as  well.


2)  Steps should be taken to  reduce algal growth wherever  possible.   The
experiments should be designed  so  that individual colonies can be  kept track
of separately.  Lighting should be continuous.

3)  Light intensity should be around 2,500 lux  with  a  standard deviation of
250 or less.

4)  Temperatures in the  test  chambers should be held close to some point
between 22°and 25°C with a standard deviation of 0.6 C or  less.  Infra-red
radiation from the fluorescent  tubes keeps the  chambers slightly above room
temperature.

5)  For most purposes a  7-day exposure is adequate,  but the first  3 days of
data are not useful because  the plants are still adjusting to their new
environment.  The glassware may be changed to reduce the  influence of epiphytic
organisms on or about the fourth  day of the  test.
                                      15

-------
6)  Toxicant concentrations  as  well  as pH,  hardness,  and alkalinity should
be measured at least  three times  in  the course of a 7-day experiment.   The
most convenient  times  are when  the fronds are added,  when the chambers are
changed to reduce periphyton growth,  and when the experiment is terminated.

7)  Aeration of  test  chambers is  not necessary.

8)  Fronds should be  counted at the  same time every day.

9)  If a notched battery jar chamber is used as described here, stainless
steel screen over the notch  is  not essential.  Any reasonably inert material
will do to hold  back  fronds  that  might escape their enclosures.  Whatever
form the chambers take,  the  table they rest on should be marked so that they
can be placed as consistently as  possible.
                                        16

-------
                                  REFERENCES
American Public Health Association.  1971.   Standard methods for the examination
     of water and wastewater.  13th ed.   New York,  N.Y.

Ashby, Eric, B. D. Bolas,  and F. T. Henderson.   1928.  Interaction of factors
     in the growth of Lemna;  I Methods  and Technique.   Ann. Bot.  42:771-782.

Ashby, Eric, and T. A. Oxley.  1935.  The interaction of factors in the growth
     of Lemna:  VI An analysis of the influence of  light intensity and
     temperature on the assimilation rate and the rate of frond multiplication.
     Ann. Bot. 49:309-336.

Blackman, G. E.  1952.  Studies in the principles of phytotoxicity.  I The
     assessment of relative toxicity. J. Exp.  Bot.  3:1-27.

Clark, N. A.  1925.  The rate of reproduction of Lemna major as a  function
     of intensity and duration of light.   J.  Phys.  Chem. 29:935-941.

DeFoe, D. L.  1975.  Multichannel Toxicant  Injection System for Flow-Through
     Bioassays.  J. Fish.  Res. Board Can.  32(4):544-546.

Eichorn, M., and H. Augsten.  1969.  Kontinuierliche Kultivierung  von Wolffia
     arrhiza (L.) Wimm. im Chemostaten.   (Continuous cultivation of Wolffia
     arrhiza (L.) Wimm. in a chemostat.  Flora,  Abt.  A,  160:576-580.

Erismann, K. H., and A. Finger.  1968.  Lemnaceen in kontinuierlicher Kultur.
     (Lemnaceae in continuous culture).   Ber. Schweiz.  Bot.  Ges. 78:5-15.

Feder, W. A., and F. Sullivan.  1969. Ozone:  depression of frond multiplication
     and floral production in duckweed.   Science 156:1373-1374.

Fekete, A., D. N. Riemer,  and H. L. Motto.   1976.  Bioassay using  common
     duckweed to evaluate  the release of available  phosphorus from pond
     sediments.  J. Aquatic Plant Manage. 14:19-25.

Fromm, F.  1946.  El Empleo de Lemna minor  L. en Ensayos Rapidos de Toxicidad.
     Cienciu.  7:214-218.

Fromm, F.  1951.  A quantitive evaluation of the Lemna test for herbicides.
     Bot. Gaz. 113:86-90.

Fromm, F.  1960  A modification of the Lemna test for phytotoxicity.  J. Agr.
     Univ. Puerto Rico 46:93-102.
                                     17

-------
Hero of Alexandria, Ca. 62 AD.  Pneumatics, Section 12.  (ed. Woodcroft, The
     pneumatics of Hero of Alexandria.  Taylor, Walton and Mabery.  London
     1851).

Hillman, W.  1961.  The Lenaceae, or duckweeds:  a review of the descriptive
     and experimental literature.  Bot. Rev.  27:221-287.

Hutner, S. H.  1953.  Comparative physiology of heterotrophic growth in plants,
     in W. E. Loomis   [ed.]  Growth and differentiation in plants.  Iowa State
     College Press, Ames, la.

Landolt, E.  1957.  Physiologische and okologische untersuchungen an
     Lemnaceen.  (Physiological and ecological investigations of the Lemnaceae)
     Ber. Schweiz. Bot. Ges. 67:272-410.

Mandiola, N. B.  1919.  Variation and selection within clonal lines of Lemna
     minor.  Genetics 41:151-182.

Mitchell, J. W., G. A. Livingston, and P. C. Marth.  1958.  Test methods with
     plant regulating chemicals.  U.S. Dep. Agr., Agr. Handb. 126.  pp. 34-36.

MDunt, D. I., and W. A. Brungs.  1967.  A simplified dosing apparatus for fish
     toxicology studies.  Water Res. 1:21-29.

Muenscher, W. C.  1944.  Aquatic plants of the United States.  Cornell
     University Press, Ithaca, N.Y.  374 p.

Parker, C.  1965.  A rapid bioassay method for the detection of herbicides
     which inhibit photosynthesis.  Weed Res. 5:181-184.

Scharer, M., C. Brunold, and K. H. Erismann.  1975.  Hemmung der Sulfataufnahme
     durch Lemna minor L. durch SC-2 in subletalen Konzentrationen.  (Inhibition
     of sulfate uptake by Lemna minor L. during aeration with sublethal
     concentrations of S02.)  Specialia:1414-1415.

Sculthorpe, C. D.  1967.  The biology of aquatic vascular plants.  Arnold,
     London.  610 p.

Strasser, R. J.  1971.  Eine einefache Anlage zur kontinuierlichen Kultivierung
     von Lemnaceen mit automatischer probeentnahme.  (Simple apparatus for
     continuous cultivation of Lemnaceae with automatic sample collection.)
     Photosynthetica 5:76-78.

Todd, G. W., J. T. Middleton, and R. F. Brewer.  1956.  Effects of air
     pollutants.  California Agr. (July):  7, 8, 14.
                                      18

-------
                                BIBLIOGRAPHY
Lemke, A. E., W. A. Brungs, and B. J. Halligan.  Manual for Construction and
     Operation of Toxicity Testing Proportional Diluters.  U.S. Environmental
     Protection Agency, Environmental Research Laboratory-Duluth, Ecological
     Research Series (in press).
                                     19

-------
                                    TECHNICAL REPORT DATA
                             (Please read Instructions on the reverse before completing)
1. REPORT NO.
   EPA-600/3-77-108
   2.
                                 3. RECIPIENT'S ACCESSION-NO.
4. TITLE AND SUBTITLE

    A FLOW-THROUGH TESTING PROCEDURE WITH DUCKWEED
    (LEMNA MINOR L.)
                                 5. REPORT DATE
                                   September  1977 issuing date
                                 6. PERFORMING ORGANIZATION CODE
7. AUTHOR(S)
                                                            8. PERFORMING ORGANIZATION REPORT NO.
    Charles T. Walbridge
9. PERFORMING ORGANIZATION NAME AND ADDRESS
    Environmental Research Laboratory  -  Duluth, MN
    Office of Research and Development
    U.S.  Environmental Protection Agency
    Duluth, Minnesota   55804
                                 10. PROGRAM ELEMENT NO.

                                    1BA608	
                                 11. CONTRACT/GRANT NO.
 12. SPONSORING AGENCY NAME AND ADDRESS

    Same as above.
                                 13. TYPE OF REPORT AND PERIOD COVERED
                                                            14. SPONSORING AGENCY CODE

                                                                EPA/600/03
 15. SUPPLEMENTARY NOTES
 16. ABSTRACT
         Lemna minor  is  one of the smallest  flowering plants.  Because of its floating
    habit,  ease of culture, and small size it  is well adapted for  laboratory investiga-
    tions.   Procedures for flow-through tests  were developed.  Testing procedures were
    developed with this  apparatus.  By using the techniques described here, the effects
    of  nutrients or toxicants, singly or in  combination, can be determined in several
    concentrations with  several replicates of  each.   Responses which  can be measured
    include changes in growth rate, changes  in death rate, changes  in timing of division
    of  colonies, color changes, changes in variability, and modification of the flowering
    response.  Emphasis  here is on changes in  growth rate determined  either by daily
    frond counts or by final frond numbers.
17.
                                 KEY WORDS AND DOCUMENT ANALYSIS
                  DESCRIPTORS
                                               b.lDENTIFIERS/OPEN ENDED TEFIMS
                                              c.  COS AT I Field/Group
   Bioassay
   Aquatic  Plants
   Plant  nutrition
   Growth regulators
   Toxicity
   Pollution
   Botany
Plant regulators
Lemna minor L.
Plant responses
Laboratory  plant
Proportional diluter
Toxicants
Nutrients
Pollutants
06 F
18. DISTRIBUTION STATEMEN1
   RELEASE TO  PUBLIC
                   19. SECURITY CLASS (ThisReport)
                     UNCLASSIFIED	
                          21. NO. OF P,
                              26
                   20. SECURITY CLASS (This page)

                     UNCLASSIFIED	
                                                                          22. PRICE
EPA Form 2220-1 (9-73)
                                                             •ft U.S. GOVERNMENT PRINTING OFFICE: 1977-757-056/6547
                                             20

-------